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Spiralia
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Spiralia
Temporal range: Cambrian–Recent[1] Possible Ediacaran fossil[2]
Scientific classification Edit this classification
Kingdom: Animalia
Subkingdom: Eumetazoa
Clade: ParaHoxozoa
Clade: Bilateria
Clade: Nephrozoa
Clade: Protostomia
Clade: Spiralia
sensu Edgecombe et al. 2011
Clades

The Spiralia are a morphologically diverse clade of protostome animals, including within their number the molluscs, annelids, platyhelminths and other taxa.[3] The term Spiralia is applied to those phyla that exhibit canonical spiral cleavage, a pattern of early development found in most members of the Lophotrochozoa.[4]

Distribution of spiralian development across phylogeny

[edit]

Members of the molluscs, annelids, platyhelminths and nemerteans have all been shown to exhibit spiral cleavage in its classical form. Other spiralian phyla (rotifers, brachiopods, phoronids, gastrotrichs, and bryozoans) are also said to display a derived form of spiral cleavage in at least a portion of their constituent species, although evidence for this is sparse.[5]

Lophotrochozoa within Spiralia

[edit]

Previously, spiral cleavage was thought to be unique to the Spiralia in the strictest sense—animals such as molluscs and annelids which exhibit classical spiral cleavage. The presence of spiral cleavage in animals such as platyhelminths could be difficult to correlate with some phylogenies.[6]

Evidence of a close relationship between molluscs, annelids and lophophorates was found in 1995 and Lophotrochozoa was defined as the group containing these taxa and all the descendants of their last common ancestor.[7] More recent research has established the Lophotrochozoa as a superphylum within the Metazoa.[8] With this understanding, the presence of spiral cleavage in polyclad platyhelminths, as well as the more traditional Spiralia, has led to the hypothesis that spiral cleavage was present ancestrally across the Lophotrochozoa as a whole.[5] With the introduction of Platytrochozoa and Rouphozoa, the cladogram is as follows, with an indication approximately how many million years ago (Mya) the clades radiated into newer clades.[9][10][11][12][13][14][15]

Protostomia
610 mya

An alternative phylogeny was given in 2019, with a basal grouping Mollusca with Entoprocta grouping named Tetraneuralia, and a second grouping of Nemertea with Platyhelminthes named Parenchymia as sister of Annelida. In their proposal and according to the original definition, Lophotrochozoa may become a senior synonym for Platytrochozoa.[1][16][17][18]

 Protostomia 


In 2019 the Rouphozoa was recovered again as a basal Platytrochozoa clade.[19]

A 2022 study supported the Trochozoa and Platyzoa hypotheses, as shown below.[20] The same year another study placed bryozoans, entoproctans and cycliophorans in the group Polyzoa as one of the earliest branches among Lophotrochozoa.[21]

Protostomia

References

[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Spiralia

View on Grokipedia
from Grokipedia
Spiralia is a major of animals within , one of the three primary lineages alongside and Deuterostomia, encompassing approximately 14 phyla of bilaterian animals and characterized by a conserved of spiral cleavage during early embryonic development. This developmental mode involves unequal, spirally arranged cell divisions that produce a determinate fate map, distinguishing Spiralia from other s with radial or modified cleavage patterns. The clade originated around the beginning of the period, representing an ancient and morphologically diverse group that includes both free-living and parasitic species across marine, freshwater, and terrestrial environments. Key phyla within Spiralia include Annelida (segmented worms such as earthworms and leeches), Mollusca (mollusks including bivalves, gastropods, and cephalopods like octopuses), Platyhelminthes (flatworms, encompassing free-living planarians and parasitic forms like schistosomes), Nemertea (ribbon worms), Rotifera (rotifers), Brachiopoda (lamp shells), Phoronida (horseshoe worms), Bryozoa (moss animals), Entoprocta, Gastrotricha, and Gnathostomulida. Many spiralian phyla exhibit trochophore larvae, a planktonic stage with a ciliary band (prototroch) used for locomotion and feeding, which is considered a synapomorphy for subgroups like Lophotrochozoa, though its presence varies across the clade. This diversity in larval forms and adult body plans—from soft-bodied invertebrates to shelled or colonial organisms—highlights Spiralia's ecological breadth, with species playing crucial roles in food webs, nutrient cycling, and human economies through fisheries, aquaculture, and biomedical research. Spiralia accounts for a significant portion, approximately 40%, of all animal phyla, making it a critical group for understanding bilaterian , genome regulation, and . Despite its vast diversity, and studies in Spiralia remain underdeveloped compared to model organisms in (e.g., and Caenorhabditis), limiting insights into conserved genetic mechanisms like clusters and ciliary band formation. Notable evolutionary novelties include spiralian-specific genes expressed in ciliary structures, which may underpin adaptations for locomotion and feeding in trochophore larvae. Ongoing phylogenomic research as of 2024 continues to refine internal relationships, such as the position of and , underscoring Spiralia's importance in reconstructing the tree of life.

Characteristics

Spiral cleavage

Spiral cleavage is a distinctive pattern of early embryonic development observed in many spiralian animals, characterized by highly stereotyped cell divisions that produce a spiral arrangement of blastomeres around the animal-vegetal axis. This mode of cleavage is determinate and , meaning cell fates are largely fixed early in development without significant regulative capacity, contrasting with the more flexible regulative development seen in other bilaterians like deuterostomes. The pattern involves oblique mitotic spindles oriented at approximately 45° to the axis, resulting in micromeres (smaller cells) forming above macromeres (larger cells) in successive tiers, typically in a counterclockwise direction from the animal pole view in most taxa, though clockwise (sinistral) arrangements occur in some groups such as certain gastropods. The process begins with the first cleavage, which produces two equal blastomeres (AB and CD), followed by the second cleavage yielding four cells where the CD pair divides unequally to form the first quartet of micromeres at the vegetal pole. The third cleavage then generates a second quartet of micromeres at the animal pole, establishing the characteristic spiral configuration with cells offset in a helical manner. Subsequent divisions continue this tiered pattern, with each quartet of micromeres contributing to specific tissues: the first quartet forms apical and oral ectoderm, the second and third quartets produce trunk ectoderm and often ectomesoderm, while the fourth quartet and underlying macromeres give rise to endoderm. A key feature is the 4d micromere, which serves as the primary mesentoblast precursor, dividing to produce mesoderm and additional endoderm in a bilateral fashion that foreshadows the adult body plan. This progression typically leads to the formation of a trochophore-like larva in many spiralian groups, such as annelids and mollusks, where the early embryo's geometry directly maps to larval structures. Variations in spiral cleavage exist across Spiralia, with the classical form—featuring equal early cleavages and strict quartet formation—prominent in annelids, mollusks, and nemerteans. In rotifers and brachiopods, the pattern is modified, often with unequal divisions or reduced spiraling, while evidence of spiralian-like cleavage is sparse and debated in gastrotrichs, potentially indicating derivation or convergence. These modifications can include shifts in cleavage direction (dextral vs. sinistral) driven by cytoskeletal asymmetries, or losses of the pattern altogether in derived lineages like cephalopods, where direct development replaces the trochophore stage. Despite these variations, the core oblique division mechanism is conserved, highlighting underlying cellular processes like ERK signaling that organize the pattern. Spiral cleavage is hypothesized as a synapomorphy of the clade, which comprises approximately 10% of known animal species, providing a developmental foundation that has been modified or lost in several subgroups, such as bryozoans and . Its evolutionary significance lies in the precise mapping it enables, which has facilitated comparative studies of bilaterian , revealing how early deterministic specification can generate bilateral from radial-like arrangements. However, the pattern's deep homology remains under investigation, with some evidence suggesting potential convergence in non-spiralian taxa or independent origins, though molecular phylogenies strongly support its role as a unifying spiralian trait.

Shared morphological traits

Certain spiralian lineages, such as those in the Tetraneuralia, exhibit a tetraneuralian characterized by four longitudinal nerve cords: paired ventral (pedal) cords and paired lateral (visceral) cords, with serotonergic perikarya and commissures. This configuration is evident in basal mollusks and entoprocts, though it is reduced to two or more cords in groups like platyhelminths. The of spiralians typically includes a formed via , where splits to create the , particularly prominent in lophotrochozoans such as annelids and mollusks. However, the ancestral spiralian condition is inferred to be acoelomate, with coelomic cavities evolving secondarily and being lost in several lineages. Larval stages often feature ciliary bands for locomotion and feeding, including the preoral prototroch and postoral metatroch. A key larval form uniting basal spiralians is the trochophore, a free-swimming, top-shaped with an apical tuft of cilia for sensory functions and prototrochal bands of compound cilia enabling swimming and particle capture. This form is ancestral for Trochozoa within Spiralia, though modified or absent in derived groups. Spiralians share specific musculature patterns, including an outer circular fiber layer, a middle diagonal layer, and an inner longitudinal fiber layer in the body wall, supporting undulatory movement. Epidermal structures commonly include a ciliated , particularly in larvae, facilitating locomotion without molting. Unlike ecdysozoans, spiralians lack , growing continuously without shedding a chitinous . Trait diversity arises through modifications, such as the secondary loss of the in , resulting in acoelomate or pseudocoelomate states that adapt to or parasitic lifestyles. These variations highlight evolutionary flexibility while retaining core spiralian features.

History

Early embryological observations

Comparative , initiated by in the 1820s through his studies of developmental stages across animals, laid the groundwork for recognizing shared patterns among , though his focus was primarily on vertebrates. Detailed observations of spiral cleavage emerged later in the . By the late , tracing advanced understanding of this cleavage mode, with researchers documenting spiral arrangements in mollusks and annelids. Edmund Beecher Wilson played a pivotal role in synthesizing these findings; in his 1892 and 1904 works, he compiled observations across multiple phyla, demonstrating that spiral cleavage produced consistent organ fates from homologous blastomeres in diverse taxa. Wilson's analyses, particularly in Experimental Studies on Germ Layers in Annelids and Mollusks (1904), emphasized the stereotyped, oblique divisions that generated quartets of cells arranged in a spiral fashion when viewed from the animal pole. These observations led to the classical grouping of "spiralian" animals, encompassing mollusks, annelids, flatworms (platyhelminths), and nemerteans, based on shared cleavage patterns that suggested potential . Early hypotheses, advanced by figures like Wilson and Edwin Conklin in the early 1900s, posited that the conserved cell lineages—such as the derivation of from the 4d blastomere—indicated a monophyletic origin for these forms, challenging earlier views of independent evolution. However, pre-molecular era interpretations treated these animals as a loose alliance rather than a formal , with ongoing debates about whether spiral cleavage represented homology or convergent to similar ecological pressures. Specific studies from the to extended these groupings by examining less-studied taxa, revealing derived variants of spiral cleavage that influenced early phylogenetic ideas. For instance, investigations into rotifers by researchers like Otto Schultze () and later compilations by Libbie Hyman (1940, drawing on data) showed a modified spiral pattern with quartet formation akin to annelids, supporting inclusion in the spiralian alliance despite reductions in cell determinacy. Similarly, entoproct embryology, documented in works by Adolf Milnes Marshall (1882) and expanded in the 1910s–1920s by Walter N. Sewell, revealed asynchronous equal spiral cleavages forming an apical rosette, though with deviations that sparked discussions on whether such patterns were primitive or secondarily modified. These findings, summarized in Walter Schleip's 1929 Die Entwicklungsweisen der Niederen Tiere, underscored the variability within spiralian development but reinforced the pattern's utility for inferring relationships among protostomes.

Establishment as a clade

The establishment of Spiralia as a monophyletic within relied heavily on molecular phylogenetic analyses beginning in the mid-1990s, which complemented earlier embryological observations of shared spiral cleavage patterns among diverse lineages. A pivotal milestone came in when Halanych et al. analyzed complete 18S ribosomal DNA sequences from lophophorate phyla (brachiopods, bryozoans, and ), demonstrating their close relationship to mollusks and annelids as protostomes, thereby proposing the to unite these groups based on shared molecular signatures in rRNA genes. This analysis resolved longstanding ambiguities in lophophorate affinities, which had been debated due to their mixture of deuterostome-like and protostome-like traits, and laid the groundwork for recognizing a broader spiralian assemblage. In the 2000s, multi-gene phylogenetic studies expanded this framework to encompass by incorporating (a proposed grouping of small-bodied, often interstitial protostomes like platyhelminths and gastrotrichs) alongside . For instance, Ruiz-Trillo et al. (2002) used heavy type II sequences to support the basal position of certain flatworms within bilaterians while reinforcing the unity of lophotrochozoan-like taxa through shared protein motifs, contributing to the emerging concept of as a comprehensive defined by molecular rather than solely morphological criteria. Subsequent multi-gene analyses, such as those employing combined nuclear and mitochondrial markers, further integrated into , highlighting convergent simplicity in body plans among these lineages rather than shared ancestry within itself. Molecular synapomorphies bolstered this clade's validity, including conserved expression patterns of NKL genes, which regulate mesodermal and neural development across spiralian taxa like annelids, mollusks, and platyhelminths. These genes, part of the NK cluster, show spiralian-specific expansions and regulatory roles that distinguish them from ecdysozoan counterparts, supporting through shared genomic architecture. Additionally, suites of microRNAs unique to spiralian lineages, such as families miR-750, miR-1000, and miR-4511, represent diagnostic genomic features absent in or Deuterostomia, with their conservation across diverse spiralians like gastrotrichs and lophotrochozoans indicating ancient origins at the clade's base. These miRNA complements, identified through , provide robust support for Spiralia's integrity by reflecting coordinated tied to spiralian developmental modes. Influential studies in the formalized Spiralia in metazoan classifications. Edgecombe et al. (2011) synthesized phylogenomic data from multiple loci, affirming Spiralia as a major alongside , with and Trochozoa as tentative internal divisions, though noting the need for denser taxon sampling to resolve boundaries. (2019) reviewed phylogenomic advancements, including large-scale datasets, which robustly confirmed Spiralia's and its position as sister to within , integrating chaetognaths into the gnathiferan via shared ortholog analyses. More recent work, such as Laumer et al. (2014) and Laumer et al. (2015), refined these boundaries using phylogenomics; for example, acanthocephalans were firmly placed within Gnathifera (a spiralian ) rather than as a separate platyzoan entity, excluding them from broader assumptions while solidifying Spiralia's scope. Debates surrounding Spiralia's establishment centered on Platyzoa's monophyly, initially proposed as a grade of simple-bodied spiralians but challenged by early molecular data showing polyphyletic origins. Transcriptomic studies resolved this uncertainty, with Laumer et al. (2015) using 402 orthologs across 90 taxa to reject Platyzoa as monophyletic—placing groups like Platyhelminthes and Gastrotricha as successive early branches within Spiralia—while affirming the clade's unity as sister to through site-heterogeneous models that accounted for compositional biases. This resolution highlighted Spiralia's ancestral non-coelomate, microscopic body plan, with Platyzoa's apparent simplicity arising from rather than shared descent. Subsequent phylogenomic and genomic studies in the , including long-read sequencing assemblies, have further confirmed Spiralia's monophyly and refined internal relationships, such as the positioning of meiofaunal lineages.

Phylogeny

Position within Protostomia

Spiralia constitutes one of the two primary clades within , the larger bilaterian group characterized by the formation of the mouth from the blastopore during early embryonic development. As the to —which encompasses arthropods, nematodes, and other molting animals—Spiralia and Ecdysozoa together form the monophyletic , excluding non-bilaterian outgroups and the basal bilaterian clade . This positioning reflects shared protostomian traits, such as determinate cleavage and , while distinguishing Protostomia from Deuterostomia, where the anus forms first from the blastopore. The placement of Spiralia as sister to is robustly supported by molecular phylogenies. Early analyses using sequences identified as a cohesive of molting , implying Spiralia (then termed ) as its complement within . Subsequent studies incorporating 28S rRNA further corroborated this bipartition. More recent phylogenomic approaches, leveraging hundreds of orthologous genes from whole-genome and data across diverse taxa, have consistently recovered high-support topologies affirming and the Spiralia- split, with bootstrap values exceeding 90% in maximum-likelihood analyses. Molecular clock estimates place the divergence of Spiralia and around 550–600 million years ago (Mya), during the late Period. The earliest unambiguous spiralian fossils appear in the early , approximately 530 Mya, including tube-like structures potentially attributable to stem-group annelids or sipunculans. Possible spiralian traces, such as simple burrows or protoconodont-like elements, extend back into the terminal (~550 Mya), though their attribution remains tentative due to limited morphological resolution. In contrast to , Spiralia lacks ecdysial molting, relying instead on non-cuticular growth mechanisms, a trait that underscores their developmental divergence within . While Spiralia shares certain developmental genes—such as subsets of Hox and ParaHox clusters—with Deuterostomia, reflecting bilaterian ancestry, its defining spiral cleavage pattern, involving oblique cell divisions, sharply distinguishes it from the radial cleavage typical of Deuterostomia.

Internal relationships and distribution of development

The internal phylogeny of Spiralia is characterized by early-branching lineages traditionally grouped under , which phylogenomic analyses show to be paraphyletic, forming a grade of small-bodied, often interstitial taxa basal to the larger . Gnathifera, including rotifers (Syndermata), gnathostomulids, and micrognathozoans (confirmed as sister to Syndermata as of 2025), emerges as the earliest diverging clade. This is followed by Rouphozoa, uniting platyhelminths (Platyhelminthes) and gastrotrichs as sister to . The position of (arrow worms) remains debated, with some analyses placing it as sister to Gnathifera within Spiralia. Within , relationships are better resolved, including clades such as Tetraneuralia ( + ) and Pleistoannelida (Annelida + ). often clusters near in proposals like Parenchymia ( + Platyhelminthes), though broader sampling is needed. Spiral cleavage is considered ancestral to Spiralia, with its characteristic oblique cell divisions and stereotyped blastomere arrangements preserved in early embryogenesis across the clade, as evidenced by comparative embryological observations in over 20 phyla. Classical spiral cleavage, featuring equal-sized blastomeres and a 45° mitotic spindle rotation, is prominently retained in Trochozoa, such as annelids (e.g., Platynereis dumerilii) and mollusks (e.g., ), where detailed tracing reveals conserved mesentoblast formation and . In basal spiralian lineages like those in the grade, the pattern is more variable: it is absent or highly modified in acoelomate flatworms, which exhibit irregular holoblastic cleavage, but persists in a derived form in gastrotrichs, with spiral-like arrangements during early divisions leading to a coeloblastula stage. These distributions are corroborated by embryological studies highlighting molecular signatures, such as Delta-Notch signaling in micromere specification, that align with phylogenetic branching. Uncertainties persist in the exact positioning of Polyzoa (bryozoans, entoprocts, and cycliophorans), which variously cluster as basal lophotrochozoans or within Tetraneuralia across datasets, potentially due to long-branch attraction in transcriptomic alignments; a 2022 study using complete gene sets supported Polyzoa as sister to . Additionally, spiral cleavage shows evidence of secondary loss in endoparasitic lineages like acanthocephalans, where direct development eliminates early cleavage patterns, reflecting adaptations to host-dependent lifestyles. Consensus trees from studies such as Laumer et al. (2019) and Marlétaz et al. (2019) depict Spiralia as a robust with these basal branches leading to , though finer resolutions await broader genomic sampling.

Classification

Lophotrochozoa

is a major clade within , proposed in 1995 by Halanych et al. based on analyses of 18S ribosomal DNA sequences that united the traditional lophophorate lineages—Brachiopoda, (also known as Ectoprocta), and Phoronida—with trochozoan groups such as and Annelida into a monophyletic assemblage. This molecular evidence revealed a close phylogenetic affinity between these disparate taxa, challenging earlier classifications that placed lophophorates nearer to deuterostomes due to shared traits like a U-shaped gut and coelomic . The clade's name reflects the defining features of its subgroups: the lophophore, a horseshoe-shaped feeding apparatus with ciliated tentacles used for suspension feeding in lophophorates, and the trochophore, a planktonic larval stage with a ciliated band for locomotion in trochozoans. The major taxa within encompass a wide array of invertebrate phyla, including (encompassing diverse classes such as bivalves like clams and oysters, gastropods like snails and slugs, and cephalopods like octopuses), Annelida (segmented worms including polychaetes, leeches, and earthworms), (ribbon worms known for their eversible proboscis), Sipuncula (peanut worms, now often classified within Annelida), (small colonial or solitary aquatic animals), Brachiopoda (lamp shells with bivalved shells), (moss animals forming encrusting colonies), and Phoronida (tube-dwelling worms). These groups exhibit key synapomorphies such as the trochophore larva in many lineages, which facilitates dispersal in marine environments; a schizocoelous mode of formation, where the body cavity develops from splits in mesodermal masses; and the ancestral retention of spiral cleavage during early embryogenesis, though modified in some derived taxa. The lophophore serves as a synapomorphy for the lophophorate subgroup, enabling efficient filter feeding in low-nutrient waters. Internally, Lophotrochozoa is divided into principal clades, with Trochozoa comprising Mollusca, Annelida, Nemertea, and associated groups like Sipuncula, characterized by trochophore larvae and often vermiform body plans adapted for burrowing or crawling. Lophophorata includes Brachiopoda, Phoronida, and Bryozoa, unified by the lophophore and supported as monophyletic in recent phylogenomic analyses using transcriptomic data. Recent refinements, such as those from phoronid genome sequencing, affirm Lophophorata monophyly while positioning Entoprocta (as Kamptozoa) outside this group, often as a sister to Bryozoa or within a broader Polyzoa clade, based on shared ciliary feeding mechanisms and molecular markers. Lophotrochozoa boasts approximately 125,000 described , the majority marine and concentrated in , which alone accounts for over 100,000 species. These organisms dominate benthic communities worldwide, serving as ecosystem engineers through activities like sediment bioturbation by annelids, shell formation by mollusks and brachiopods that provide habitats, and suspension feeding by bryozoans and that regulate nutrients. Their ecological roles extend to support, with many taxa prey for higher predators and contributors to carbon cycling in seafloor sediments.

Platyzoa and other subgroups

Platyzoa is a proposed within Spiralia, first defined by Thomas Cavalier-Smith in 1998 to encompass a diverse assemblage of mostly acoelomate or pseudocoelomate animals characterized by flattened, simple, or worm-like body plans. This grouping includes several phyla that were historically considered basal or transitional in bilaterian evolution due to their lack of a and reduced organ systems. The clade was motivated by shared morphological traits such as dorsoventrally flattened bodies and or parasitic lifestyles, though molecular data have since challenged its . The major taxa within Platyzoa include Platyhelminthes (flatworms), which comprise free-living and parasitic forms with dorsoventrally flattened bodies; Gastrotricha, microscopic interstitial animals with spiny cuticles; Rotifera (wheel animals), known for their ciliated corona used in locomotion and feeding; Acanthocephala (thorny-headed worms), obligate endoparasites with a armed with hooks; Gnathostomulida, tiny jawed worms inhabiting marine sands; and Micrognathozoa, a rare with a single described featuring complex jaw structures. These groups collectively represent thousands of , predominantly microscopic in size and adapted to freshwater, marine, or parasitic niches, often as part of the meiofaunal communities in sediments or host tissues. Key features across Platyzoa include simplified body organizations without segmentation, with many taxa exhibiting modified or reduced versions of spiral cleavage—such as duet-spiral patterns in polyclad flatworms or its complete loss in some parasitic platyhelminths—reflecting adaptations to their environments. Rotifers, for instance, display a distinctive moniliform (bead-like) body segmentation and a retractable corona, aiding in their pseudocoelomate structure. Debated inclusions in have included (encompassing Dicyemida and ), small vermiform parasites of once thought to represent primitive bilaterians. Recent phylogenomic analyses, such as a 2022 study using multiple methods, support the of within a broader monophyletic sensu lato, often as sister to or Platyhelminthes, though some Bayesian analyses indicate . In some earlier phylogenies, (an alternative name for or related ectoprocts) was considered a potential or separate clade within broader Spiralia, though modern studies relegate it firmly to . Phylogenetic challenges persist for , with early molecular support giving way to evidence of in some 2014 phylogenomic data (e.g., Struck et al.), where groups like Platyhelminthes and Gastrotricha do not form a cohesive but disperse across basal Spiralia positions, likely due to long-branch attraction artifacts. However, further analyses as of 2022 often recover of s.l. in maximum-likelihood and coalescent-based phylogenomics, with Gnathifera as basal and included, suggesting shared ancestry for simple body plans rather than pure convergence, though debate continues with mixed results from different methods. Ecologically, taxa dominate as in freshwater and marine habitats, filling roles in cycling and as intermediate hosts in parasitic food webs, underscoring their importance despite phylogenetic uncertainties.

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